Transcript profiling of a novel plant meristem, the monocot cambium.

While monocots lack the ability to produce a vascular cambium or woody growth, some monocot lineages evolved a novel lateral meristem, the monocot cambium, which supports secondary radial growth of stems. In contrast to the vascular cambium found in woody angiosperm and gymnosperm species, the monocot cambium produces secondary vascular bundles, which have an amphivasal organization of tracheids encircling a central strand of phloem. Currently there is no information concerning the molecular genetic basis of the development or evolution of the monocot cambium. Here we report high-quality transcriptomes for monocot cambium and early derivative tissues in two monocot genera, Yucca and Cordyline. Monocot cambium transcript profiles were compared to those of vascular cambia and secondary xylem tissues of two forest tree species, Populus trichocarpa and Eucalyptus grandis. Monocot cambium transcript levels showed that there are extensive overlaps between the regulation of monocot cambia and vascular cambia. Candidate regulatory genes that vary between the monocot and vascular cambia were also identified, and included members of the KANADI and CLE families involved in polarity and cell-cell signaling, respectively. We suggest that the monocot cambium may have evolved in part through reactivation of genetic mechanisms involved in vascular cambium regulation.

Floral scent of Joshua trees (Yucca brevifolia sensu lato): Divergence in scent profiles between species but breakdown of signal integrity in a narrow hybrid zone.

PREMISE OF THE STUDY: The role of floral scent in facilitating reproductive isolation between closely related plants remains poorly understood. Yucca brevifolia and Yucca jaegeriana are pollinated by different moth species in allopatry, but in a narrow contact zone, pollinator-host specificity breaks down, resulting in hybridization between species. We explored the chemical basis for reproductive isolation and hybridization in these Joshua trees by characterizing the floral scent of each species in allopatry, analyzing scent profiles from trees in the contact zone, and matching these data with genotypic and phenotypic data. METHODS: We analyzed floral volatiles using gas chromatography-mass spectrometry, tested for species divergence of scent profiles and classified trees in the contact zone as hybrid or either parental species. We used floral and vegetative morphological data and genotypic data to classify trees and analyzed whether certain trait combinations were more correlated than others with respect to assignment of trees and whether frequencies of classified tree types differed depending on which data set was used. KEY RESULTS: The Joshua tree floral scent included oxygenated 8-carbon compounds not reported for other yuccas. The two species differed (P < 0.001) in scent profiles. In the contact zone, many hybrids were found, and phenotypic traits were generally weakly correlated, which may be explained by extensive gene flow between species or by exposure to different selection pressures. CONCLUSIONS: Although the two Joshua tree species produce distinct floral scent profiles, it is insufficient to prevent attraction of associated pollinators to both hosts. Instead, floral morphology may be the key trait mediating gene flow between species.

Gas exchange and leaf anatomy of a C3-CAM hybrid, Yucca gloriosa (Asparagaceae).

While the majority of plants use the typical C3 carbon metabolic pathway, ~6% of angiosperms have adapted to carbon limitation as a result of water stress by employing a modified form of photosynthesis known as Crassulacean acid metabolism (CAM). CAM plants concentrate carbon in the cells by temporally separating atmospheric carbon acquisition from fixation into carbohydrates. CAM has been studied for decades, but the evolutionary progression from C3 to CAM remains obscure. In order to better understand the morphological and physiological characteristics associated with CAM photosynthesis, phenotypic variation was assessed in Yucca aloifolia, a CAM species, Yucca filamentosa, a C3 species, and Yucca gloriosa, a hybrid species derived from these two yuccas exhibiting intermediate C3-CAM characteristics. Gas exchange, titratable leaf acidity, and leaf anatomical traits of all three species were assayed in a common garden under well-watered and drought-stressed conditions. Yucca gloriosa showed intermediate phenotypes for nearly all traits measured, including the ability to acquire carbon at night. Using the variation found among individuals of all three species, correlations between traits were assessed to better understand how leaf anatomy and CAM physiology are related. Yucca gloriosa may be constrained by a number of traits which prevent it from using CAM to as high a degree as Y. aloifolia. The intermediate nature of Y. gloriosa makes it a promising system in which to study the evolution of CAM.

Absence of population-level phenotype matching in an obligate pollination mutualism.

Coevolution is thought to promote evolutionary change between demes that ultimately results in speciation. If this is the case, then we should expect to see similar patterns of trait matching and phenotypic divergence between populations and between species in model systems for coevolution. As measures of divergence are frequently only available at one scale (population level or taxon level), this contention is rarely tested directly. Here, we use the case of co-divergence between different varieties of Joshua tree Yucca brevifolia (Agavaceae) and their obligate pollinators, two yucca moths (Tegeticula spp. Prodoxidae), to test for trait matching between taxa and among populations. Using model selection, we show that there is trait matching between mutualists at the taxon level, but once we account for differences between taxa, there is no indication of trait matching in local populations. This result differs from similar studies in other coevolving systems. We hypothesize that this discrepancy arises because coevolution in obligate mutualisms favours divergence less strongly than coevolution in other systems, such as host­parasite interactions.

Coevolution and divergence in the Joshua tree/yucca moth mutualism.

Theory suggests that coevolution drives diversification in obligate pollination mutualism, but it has been difficult to disentangle the effects of coevolution from other factors. We test the hypothesis that differential selection by two sister species of pollinating yucca moths (Tegeticula spp.) drove divergence between two varieties of the Joshua tree (Yucca brevifolia) by comparing measures of differentiation in floral and vegetative features. We show that floral features associated with pollination evolved more rapidly than vegetative features extrinsic to the interaction and that a key floral feature involved in the mutualism is more differentiated than any other and matches equivalent differences in the morphology of the pollinating moths. A phylogenetically based, ancestral states reconstruction shows that differences in moth morphology arose in the time since they first became associated with Joshua trees. These results suggest that coevolution, rather than extrinsic environmental factors, has driven divergence in this obligate pollination mutualism.